8 Photosynthesis: Energy from the Sun

Powered by sunlight, green plants convert CO2 and water into car-
bohydrates by a process called photosynthesis. The emergence of
this metabolic pathway was a key event in the evolution of life.
Photosynthesizing organisms, called autotrophs (“self-feeders”), use
solar energy to make their own food from simple chemicals in the
environment. In this way, they provide an entry point to the biosphere for chemical
energy. Heterotrophs (“other-feeders”) cannot photosynthesize, and they depend on
autotrophs (or other heterotrophs) for the raw materials of metabolism, such as glu-
cose.
The “food chain” from autotrophs to heterotrophs requires a lot of photosynthe-
sis. On the African plain, it takes 6 acres of grassland to convert enough CO2 into
plant matter to support the growth of one gazelle that consumes the grass. Globally,
more than 10 billion tons of carbon is fixed—converted from being part of a simple
gas (CO2) into a more complex molecule (carbohydrate)—by plants every year. This
huge amount of photosynthetically-fixed carbon is available for use by all species
that need it.
Humans consume a huge amount of Earth’s photosynthetic output. Recent calcu-
lations of total plant growth in agriculture, pastures, and forests and the products con-
sumed by people indicate that one-third of all the carbon fixed annually is appropri- Primary Producers Plants, through
photosynthesis, are the basis of life on
ated by humans, leaving two-thirds for the entire remainder of the biosphere. This is Earth. The energy stored in these soybeans
by far the greatest proportion of consumption for any single species in known history, being cultivated on a farm in Kansas will be
Is this situation sustainable? Conferences harvested and used by human beings.
such as the 2002 United Nations Confer-
ence on Sustainability have demonstrated
concern for our photosynthetic future.
An important first step in examining
ecological sustainability is a thorough
understanding of photosynthesis. The
process of photosynthesis can be neatly
broken down into two steps. The first step
is the conversion of energy from light to
chemical bonds in reduced electron carri-
ers and ATP. In the second step, these two
sources of chemical energy are used to
drive the synthesis of carbohydrates from
carbon dioxide. In this chapter, we will ex-
amine these two processes and show how
they are related to each other and to plant
growth.
146 CHAPTER EIGHT

How do the carbons become linked to form sugars? And

Identifying Photosynthetic Reactants
and Products
By the beginning of the nineteenth century, scientists under-
stood the broad outlines of photosynthesis. It was known to
use three principal ingredients—water, carbon dioxide (CO2),
and light—and to produce not only carbohydrates but also
oxygen gas (O2). Scientists had learned several things:

The water for photosynthesis in land plants comes pri-
marily from the soil and must travel from the roots to the
leaves.

Carbon dioxide is taken in, and water and O2 are re-
leased, through tiny openings in leaves, called stomata
(singular, stoma) (Figure 8.1).

Light is absolutely necessary for the production of oxy-
gen and carbohydrates.
By 1804, scientists could summarize photosynthesis as follows:
carbon dioxide + water + light energy → sugar + oxygen
which turns into an equation that is the reverse of the overall
equation for cellular respiration given in Chapter 7:
6 CO2 + 6 H2O → C6H12O6 + 6 O2
Although correct, these statements say nothing about the de-
tails of the process, and in fact, in detail, photosynthesis is not
the reverse of cellular respiration. What are the reactions of
photosynthesis? What role does light play in these reactions?
does the oxygen gas come from the CO2 or from the H2O?
Almost a century and a half passed before the source of
the O2 released during photosynthesis was determined. One
of the first uses of an isotopic tracer in biological research re-
sulted in its identification. In these experiments, two groups
of green plants were allowed to carry on photosynthesis.
Plants in the first group were supplied with water contain-
ing the oxygen isotope 18O and with CO2 containing only the
common oxygen isotope 16O; plants in the second group
were supplied with CO2 labeled with 18O and water con-
taining only 16O.
When oxygen gas was collected from each group of plants
and analyzed, it was found that O2 containing 18O was pro-
duced in abundance by the plants that had been given 18O-
labeled water, but not by the plants given 18O-labeled CO2.
These results showed that all the oxygen gas produced dur-
ing photosynthesis comes from water (Figure 8.2). This dis-
covery is reflected in a revised balanced equation:
6 CO2 + 12 H2O → C6H12O6 + 6 O2 + 6 H2O
Water appears on both sides of the equation because water
is both used as a reactant (the twelve molecules on the left)
and released as a product (the six new ones on the right). In
this revised equation, there are now sufficient water mole-
cules to account for all the oxygen gas produced.

EXPERIMENT
Question: What is the source of the O2 produced by photosynthesis?

H2O Experiment 1 Experiment 2

Sunlight
Carbon dioxide enters and O2
O2 exits the leaves through H2 18O , CO2 H2O, C 18O2
CO2 openings on the leaf surface METHOD
called stomata. These pores
can be open or closed. Plants were Plants were given
given isotope- isotope-labeled
labeled water carbon dioxide
(H218O), and (C18O2), and
unlabeled CO2. unlabeled water.
Leaf
Stem Sugars, the products of
photosynthesis, are
transported throughout
the plant body. 18O
2
O2

The oxygen The oxygen

RESULTS released released was
was labeled. unlabeled.
Root
H2O
Conclusion: Water is the source of the O2 produced by photosynthesis.

8.2 Water Is the Source of the Oxygen Produced by

8.1 The Ingredients for Photosynthesis A typical terrestrial plant Photosynthesis Because only plants given isotope-labeled
uses light from the sun, water from the soil, and carbon dioxide from water released labeled O2, this experiment showed that water
the atmosphere to form organic compounds by photosynthesis. is the source of the oxygen released during photosynthesis.
 PHOTOSYNTHESIS: ENERGY FROM THE SUN 147

The two pathways are linked by the exchange of ATP and

The Two Pathways of Photosynthesis:
An Overview
The equation above summarizes the overall process of pho-
tosynthesis. However, like glycolysis and the other metabolic
pathways that yield energy in cells, photosynthesis consists
not of one single reaction but of many reactions. The reac-
tions of photosynthesis can be divided into two pathways:

The light reactions are driven by light energy. This
pathway produces ATP and a reduced electron carrier
(NADPH + H+).

The second pathway, called the Calvin–Benson cycle,
does not use light directly. It uses ATP, NADPH + H+
(made by the light reactions), and CO2 to produce
sugars.
The reactions of the Calvin–Benson cycle are sometimes
called the dark reactions because they do not directly require
light energy. However, both pathways stop in the dark be-
cause ATP synthesis and NADP+ reduction require light. The
reactions of both pathways proceed within the chloroplast,
but they reside in different parts of that organelle (Figure 8.3).
Stroma
ADP, and of NADP+ and NADPH, and the rate of each set of
reactions depends on the rate of the other.
We will discuss the light reactions at length, to be followed
by the details of the Calvin–Benson cycle. However, because
these two photosynthetic pathways are powered by the en-
ergy of light, we begin by discussing the physical nature of
light and the nature of the specific photosynthetic molecules
that capture its energy.
The Interactions of Light and Pigments
Light is a source of both energy and information. In later
chapters, we’ll examine the many roles of light in the trans-
mission of information. In this chapter, our focus is on light
as a source of energy.
Light behaves as both a particle and a wave
Light is a form of electromagnetic radiation. It comes in dis-
crete packets called photons. Light also behaves as if it were
propagated in waves. The amount of energy contained in a
single photon is inversely proportional to its wavelength: the
shorter the wavelength, the greater the energy of the photons.
For example, a photon of red light of wavelength 660 nm has
less energy than a photon of blue light at 430 nm.

Thylakoids

Light
(photon)

ELECTRON
TRANSPORT
Chloroplast
Thylakoid
Chlorophyll

H2O O2 Light
reactions Thylakoid

ATP NADPH
CYCLE CYCLE
ADP + Pi ATP NADP+ NADPH + H+
Stroma 8.3 An Overview of
Photosynthesis The com-
plete photosynthetic reac-
CO2 fixation tion comprises two path-
CALVIN–
CO2 reactions ways: the light reactions
BENSON Sugars
and the Calvin–Benson
CYCLE
cycle. These reactions take
place in the thylakoids
(shown in the micrograph
at right) and stroma of
chloroplasts, respectively.
148 CHAPTER EIGHT

Two things are required for photons to be active in a bio-
logical process:

Photons must be absorbed by a receptive molecule.

Photons must have sufficient energy to perform the
chemical work required.
Absorbing a photon puts a pigment in an excited state
When a photon meets a molecule, one of three things happens:

The photon may bounce off the molecule—it may be
scattered.

The photon may pass through the molecule—it may be
transmitted.

The photon may be absorbed by the molecule.
however, cannot disappear, because energy is neither created
nor destroyed.
When a molecule absorbs a photon, that molecule acquires
the energy of the photon. It is thereby raised from a ground
state (lower energy) to an excited state (higher energy) (Fig-
ure 8.4a). The difference in energy between the molecule’s ex-
cited state and its ground state is exactly equal to the energy
of the absorbed photon. The increase in energy boosts one of
the electrons within the molecule into an orbital farther from
its nucleus; this electron is now held less firmly (Figure 8.4b),
making the molecule more chemically reactive, as we will see
later in the chapter.
The electromagnetic spectrum (Figure 8.5) shows the wide
range of wavelengths (and hence, energy levels) that photons
can have. The specific wavelengths absorbed by a particular

Neither of the first two outcomes causes any change in the

molecule. In the third case, the photon disappears. Its energy, Cosmic rays
Gamma rays

(a)
Excited Wavelength (nm)
state
1
X rays
Increasing energy

Shorter wavelengths are

Absorption more energetic.
Photon of photon by
molecule 10

400
Ground Violet
state
Ultraviolet (UV) 10 2 Blue

When a molecule in the ground state absorbs a photon,

it is raised to an excited state and possesses more energy. 500 Green
Visible
light Yellow
10 3
(b) Photon
Orange
Electron 600
Nucleus

10 4 Red

700
Infrared (IR)

10 5

Ground state Excited state Longer wavelengths are

less energetic.

The absorption of the photon boosts an electron to a

shell farther from its atomic nucleus. 10 6

Microwaves
8.4 Exciting a Molecule (a) When a molecule absorbs the energy Radio waves
of a photon, it is raised from a ground state to an excited state. (b) In
the excited state, an electron is boosted to a more distant shell, 8.5 The Electromagnetic Spectrum The portion of the electromag-
where it is held less firmly. netic spectrum that is visible to humans is shown in detail at the right.
 PHOTOSYNTHESIS: ENERGY FROM THE SUN 149

molecule are characteristic of that type of molecule. Mole-
cules that absorb wavelengths in the visible spectrum—that re-
gion of the spectrum that is visible to humans—are called
pigments.
When a beam of white light (light containing visible light
of all wavelengths) falls on a pigment, certain wavelengths
of the light are absorbed. The remaining wavelengths, which
are scattered or transmitted, make the pigment appear to us
to be colored. For example, if a pigment absorbs both blue
and red light—as chlorophyll does—what we see is the re-
maining light, which is primarily green.
Absorbed wavelengths correlate with biological activity
If we plot the wavelengths of the light absorbed by a puri-
fied molecule, the result is an absorption spectrum for that
molecule. If we plot the biological activity of a photosynthetic
organism as a function of the wavelengths of light to which
the organism is exposed, the result is an action spectrum.
Figure 8.6 shows the absorption spectrum for a pigment,
Blue and orange-red wavelengths cause
the highest rates of photosynthesis.
(a)
Absorption by pigment
chlorophyll a, isolated from the leaves of a plant and the ac-
tion spectrum for photosynthetic activity for the same plant.
A comparison of the two spectra shows that the wavelengths
at which photosynthesis is maximal are the same wave-
lengths at which chlorophyll a absorbs light.
Photosynthesis uses energy absorbed
by several pigments
The light energy used for photosynthesis is not absorbed by
just a single type of pigment. Instead, several different pig-
ments with different absorption spectra absorb the energy
that is eventually used for photosynthesis. In photosynthetic
organisms of all kinds (plants, protists, and bacteria), these
pigments include chlorophylls, carotenoids, and phycobilins.
In plants, two chlorophylls predominate: chlorophyll a
and chlorophyll b. These two molecules differ only slightly
in their molecular structure. Both have a complex ring struc-
ture similar to that of the heme group of hemoglobin. In the
center of each chlorophyll ring is a magnesium atom, and at-
tached at a peripheral location on the ring is a long hydro-
carbon “tail,” which can adhere the chlorophyll molecule to
proteins in the hydrophobic portion of the thylakoid mem-
brane (Figure 8.7).
We saw in Figure 8.6 that the chlorophylls absorb blue and
red wavelengths, which are near the two ends of the visible
spectrum. Thus, if only chlorophyll pigments were active in
photosynthesis, much of the visible spectrum would go un-
used. However, all photosynthetic organisms possess acces-

Absorbtion spectrum sory pigments, which absorb photons intermediate in energy

of chlorophyll a between the red and the blue wavelengths (for instance, yel-
low light) and then transfer a portion of that energy to the
chlorophylls.
Among these accessory pigments are carotenoids, such as
β-carotene (see Figure 3.22), which absorb photons in the blue
and blue-green wavelengths and appear deep yellow. The
phycobilins, which are found in red algae and in cyanobacte-
(b)
ria, absorb various yellow-green, yellow, and orange wave-
lengths. Such accessory pigments, in collaboration with the
Action spectrum chlorophylls, constitute an energy-absorbing system cover-
of Anacharis
ing much of the visible spectrum.
Photosynthesis by plant

Light absorption results in photochemical change

After a pigment molecule absorbs a photon and enters an ex-
cited state (see Figure 8.4), that molecule may return to the
ground state. When this happens, some of the absorbed en-

400 450 500 550 600 650 700 750 8.6 Absorption and Action Spectra The absorption spectrum (a) of the puri-
Wavelength (nm) fied pigment chlorophyll a from the aquatic plant Anacharis is similar to the
Visible spectrum action spectrum (b) obtained when different wavelengths of light are shone on
the intact plant and the rate of photosynthesis is measured (b).
150 CHAPTER EIGHT

CH2 8.7 The Molecular Structure of Chlorophyll Chlorophyll consists of a

(CHO in chlorophyll b)
CH CH3
complex ring structure (shaded area) with a magnesium atom at the center,
CH3
plus a hydrocarbon “tail.”The “tail” anchors chlorophyll molecules to the thy-
CH
lakoid membrane. Chlorophyll a and chlorophyll b are identical except for the
H3 C CH2
Light is absorbed by the replacement of a methyl group (—CH3) with an aldehyde group (—CHO) at
N N complex ring structure of the upper right.
HC CH
a chlorophyll molecule.
Mg

N N
H 3C
CH3 Chlorophyll Stroma charged. In plants, the pigment molecule in the re-
H molecules action center is always a molecule of chlorophyll a.
H
H2C
H C C
There are many other chlorophyll a molecules in the
CH2 C O O antenna system, but all of them absorb light at
O C O shorter wavelengths than does the molecule in the
CH3 reaction center.
O

CH2

CH
Proteins
Thylakoid Excited chlorophyll in the reaction center acts
membrane
Hydrocarbon tails secure as a reducing agent for electron transport
chlorophyll molecules to Thylakoid interior
hydrophobic proteins inside
Ultimately, photosynthesis stores chemical energy
the thylakoid membrane. by using the excited chlorophyll molecule in the re-
action center as a reducing agent to reduce a stable
electron acceptor (Figure 8.8). Ground-state chlorophyll (sym-
bolized Chl) is not much of a reducing agent, but excited
chlorophyll (Chl*) is a good one. To understand the reducing
capability of Chl*, recall that in an excited molecule, one of
ergy is given off as heat and the rest is given off as light en- the electrons is zipping around in an orbital farther away
ergy, or fluorescence. Because some of the absorbed light en- from its nucleus. Less tightly held, this electron can be passed
ergy is lost as heat, the fluorescence has less energy and
longer wavelengths than the absorbed light. When there is
fluorescence, there are no permanent chemical changes or bi- Excited state
ological functions—no chemical work is done. The energized
electron from
Any pigment molecule can become excited when its ab- e–
the chlorophyll
e–
sorption spectrum matches the energies of incoming photons. molecules is
passed on to an
If fluorescence does not occur, that pigment molecule may Electron electron acceptor.
Photon
pass the absorbed energy along to another molecule, provided acceptor
that the target molecule is very near, has the right orientation,
Reaction center
and has the appropriate structure to receive the energy. (chlorophyll molecule)
The pigments in photosynthetic organisms are arranged
into energy-absorbing antenna systems. In these systems, the
pigments are packed together and attached to thylakoid
Chlorophyll
membrane proteins in such a way that the excitation energy molecules
from an absorbed photon can be passed along from one pig-
Hydrocarbon
ment molecule in the system to another (see Figures 8.7 and tails
8.8). Excitation energy moves from pigments that absorb
Proteins
shorter wavelengths (higher energy) to pigments that absorb
Antenna system embedded
longer wavelengths (lower energy). Thus the excitation ends in thylakoid membrane
up in the one pigment molecule in the antenna system that
8.8 Energy Transfer and Electron Transport Rather than being
absorbs the longest wavelengths; this molecule is in the re-
lost as fluorescence, energy from a photon may be transferred from
action center of the antenna system. one pigment molecule to another, preserving the energy for bio-
It is the reaction center that converts the light absorbed chemical work. In an antenna system, an excited pigment molecule
into chemical energy. It is in the reaction center that a mole- can transfer energy through a series of other pigment molecules to a
chlorophyll molecule in the reaction center. That molecule may
cule absorbs sufficient energy that it actually gives up its ex- become sufficiently excited that it gives up its excited electron, which
cited electron (is chemically oxidized) and becomes positively can then be passed on to an electron carrier.
 PHOTOSYNTHESIS: ENERGY FROM THE SUN 151

on in a redox reaction to an oxidizing agent. Thus Chl* (but
not Chl) can react with an oxidizing agent A in a reaction like
this:
Chl* + A → Chl+ + A–
This, then, is the first consequence of light absorption by
chlorophyll: The chlorophyll becomes a reducing agent and
participates in a redox reaction.
As we are about to see, the further adventures of the elec-
trons from chlorophyll reduce the electron carrier NADP+
and generate a proton-motive force that is eventually used to
synthesize ATP.
ipates in catabolism, NADP+ is used in anabolic (synthetic)
reactions, such as carbohydrate synthesis from CO2, that re-
quire energy from reducing power.
Electron transport in the thylakoid membrane sets up a
charge separation, just as electron transport in the inner mi-
tochondrial membrane does (see Chapter 7). This potential
energy is captured by the chemiosmotic synthesis of ATP in a
process called photophosphorylation.
Both NADPH + H+ and ATP are used in the Calvin–Ben-
son cycle as a source of energy for the endergonic synthesis
of carbohydrates (see Figure 8.3).
There are two different systems of electron transport in
photosynthesis:

The Light Reactions: Electron Transport,
Reductions, and Photophosphorylation
The energized electron that leaves the activated chlorophyll
in the reaction center needs somewhere to go. It immediately
participates in a series of oxidation-reduction (redox) reac-
tions. The energy-rich electron is passed through a chain of
electron carriers in the thylakoid membrane in a process
termed electron transport. Two energy-rich products of the
light reactions, NADPH + H+ and ATP, are the result.
The energy-rich NADPH + H+ is a stable, reduced coen-
zyme. Its oxidized form is NADP+ (nicotinamide adenine
dinucleotide phosphate). Just as NAD+ couples the meta-
bolic pathways of cellular respiration, NADP+ couples the
two photosynthetic pathways. NADP+ is identical to NAD+
except that the former has an additional phosphate group at-
tached to each ribose (see Figure 7.4). Whereas NAD+ partic-

Noncyclic electron transport produces NADPH + H+
and ATP.

Cyclic electron transport produces only ATP.
We’ll consider these two systems before considering the role
of chemiosmosis in phosphorylation—a process that is very
similar to oxidative phosphorylation in mitochondria (see
Chapter 7).
Noncyclic electron transport produces ATP and NADPH
In noncyclic electron transport, light energy is used to oxi-
dize water, forming O2, H+, and electrons. Follow the steps
in Figure 8.9 as you read this section.
8.9 Noncyclic Electron Transport Uses Two Photosystems
Photosystems I and II both make use of the excited chlorophyll mole-
cules of their respective reaction centers.

Photosystem II Electron transport chain Photosystem I Fd

e– NADP+
e– e– reductase
e–
Energy of molecules

NADP+
Photon NADPH
H2O +
H+ H+ + H+
Photon e–
O2 + 2 H+ e– P700
ADP + Pi ATP
P680

1 Photosystem II uses
light to oxidize water
molecules, producing
electrons, H+, and O2.
2 The Chl molecule in the
reaction center of
photosystem II absorbs
light maximally at
680 nm, becoming Chl*.
3 H+ from H2O and electron 4 The Chl molecule in the 5 Photosystem I reduces
transport through the reaction center of an oxidizing agent,
redox chain is captured photosystem I absorbs ferredoxin (Fd), which
for the chemiosmotic light maximally at 700 nm, in turn reduces NADP+
synthesis of ATP. becoming Chl*. to NADPH + H+.
152 CHAPTER EIGHT

Electrons from water replenish the electrons that chloro-
phyll molecules lose when they are excited by light. As the
electrons are passed from water to chlorophyll, and ulti-
mately to NADP+, they pass through a chain of electron car-
riers. These redox reactions are exergonic, and some of the
free energy released is used ultimately to form ATP by a
chemiosmotic mechanism.
TWO PHOTOSYSTEMS ARE REQUIRED. Noncyclic electron trans-
port requires the participation of two different photosystems.
These photosystems are light-driven molecular units, each
of which consists of many chlorophyll molecules and acces-
sory pigments bound to proteins in separate energy-absorb-
ing antenna systems.

Photosystem I uses light energy to reduce NADP+ to
NADPH + H+.

Photosystem II uses light energy to oxidize water mole-
cules, producing electrons, protons (H+), and O2.
P700*, which then leads to the reduction of an oxidizing agent
called ferredoxin (Fd) and the production of P700+. Then P700+
returns to the ground state by accepting electrons passed
through the redox chain from photosystem II.
With this accounting for the source of the electrons enter-
ing photosystem II, we can now consider the fate of the elec-
trons from photosystem I. These electrons are used in the last
step of noncyclic electron transport, in which two electrons
and two protons are used to reduce a molecule of NADP+ to
NADPH + H+.
In summary:

Noncyclic electron transport uses a molecule of water,
four photons (two each absorbed by photosystems I and
II), one molecule each of NADP+ and ADP, and one Pi.

Noncyclic electron transport produces NADPH + H+
and ATP and half a molecule of oxygen ( 1/2 O2).

The reaction center for photosystem I contains a chloro-
phyll a molecule called P700 because it can best absorb light of
wavelength 700 nm. The reaction center for photosystem II
contains a chlorophyll a molecule called P680 because it ab-
sorbs light maximally at 680 nm. Thus photosystem II re-
quires photons that are somewhat more energetic (i.e., shorter
wavelengths) than those required by photosystem I. To keep
noncyclic electron transport going, both photosystems I and
II must constantly be absorbing light, thereby boosting elec-
trons to higher orbitals from which they may be captured by
specific oxidizing agents.
Cyclic electron transport produces ATP but no NADPH
Noncyclic electron transport produces ATP and NADPH +
H+. However, as we will see, the Calvin–Benson cycle uses
more ATP than NADPH + H+. Cyclic electron transport oc-
curs in some organisms when the ratio of NADPH + H+ to
NADP+ in the chloroplast is high. This process, which pro-
duces only ATP, is called cyclic because an electron passed
from an excited chlorophyll molecule at the outset cycles back
to the same chlorophyll molecule at the end of the chain of reac-
tions (Figure 8.10).

DETAILS OF THE REACTIONS. The reac- 1 In cyclic electron flow, excited electron 2 Reduced ferredoxin then
tions of noncyclic electron transport transport and chlorophylls pass electrons reduces plastoquinone,
to an oxidizing agent, ferredoxin, leaving and so forth, down the
from water to NADP+ are depicted positively charged chlorophyll (Chl+). redox chain from
in Figure 8.9. Photosystem II absorbs ferredoxin through
photons, sending electrons from P680 plastocyanin.
Photosystem I Electron transport chain
to the primary electron acceptor—
the first carrier in the redox chain— e–
e–
and causing P680 to become oxidized 3 Energy from
Energy of molecules

to P680+. Electrons from the oxidation electron flow is

of water are passed to P680+, reducing captured for
chemiosmotic
it once again to P680, which can then synthesis of ATP.
absorb more photons. The electron Photon
from photosystem II passes through
ADP + Pi ATP
a series of exergonic reactions in the P700
e–
4 The last reduced
redox chain that are indirectly cou- electron carrier
pled across the thylakoid membrane (plastocyanin) passes
to proton pumping. This pumping electrons to electron-
deficient chlorophyll,
creates a proton gradient that pro- allowing the reactions
duces energy for ATP synthesis. 8.10 Cyclic Electron Transport Traps Light Energy as ATP Cyclic to start again.
electron transport produces ATP, but no NADPH + H+. The same
In photosystem I, the reaction cen- chlorophyll molecule passes on the electrons that start the reactions
ter containing P700 becomes excited to and receives the electrons at the end to start the process over again.
 PHOTOSYNTHESIS: ENERGY FROM THE SUN 153

Before cyclic electron transport begins, P700, the reaction
center chlorophyll of photosystem I, is in the ground state. It
absorbs a photon and becomes P700*. The P700* then reacts
with oxidized ferredoxin (Fdox) to produce reduced ferre-
doxin (Fdred). The reaction is exergonic, releasing free energy.
Reduced ferredoxin (Fdred) passes its added electron to a dif-
ferent oxidizing agent, plastoquinone (PQ, a small organic mol-
ecule), which pumps 2 H+ back across the thylakoid mem-
brane. Thus, Fdred reduces PQ, and PQred passes the electron
to a cytochrome complex (Cyt). The electron continues down
the electron transport chain until it completes its cycle by re-
turning to P700+, resulting in a restoration of its uncharged
form, P700. By the time the electron from P700* travels through
ELECTRON
TRANSPORT
the redox chain by way of plastocyanin (PC), and comes back
to reduce P700+, all the energy from the original photon has
been released. This cycle is a series of redox reactions, each
exergonic, and the released energy is stored in the form of a
proton gradient that can be used to produce ATP.
Having seen how a proton gradient is established across
the thylakoid membrane, we’ll now examine in more detail
the role of this gradient in ATP synthesis.
Chemiosmosis is the source of the ATP
produced in photophosphorylation
In Chapter 7 we considered the chemiosmotic mechanism for
ATP formation in the mitochondrion. The chemiosmotic
mechanism also operates in photophosphorylation (Figure
8.11). In chloroplasts, as in mitochondria, electron transport
through the redox chain is coupled to the transport of pro-

ATP NADPH 8.11 Chloroplasts Form ATP Chemiosmotically

CYCLE CYCLE
Protons (H+) pumped across the thylakoid membrane
from the stroma during electron transport make the
CALVIN–
BENSON interior of the thylakoid more acidic than the stroma.
CYCLE
Thylakoid
Driven by this pH difference, the protons diffuse back to the
interior stroma through ATP synthase channels, which couple the ener-
gy of proton diffusion to the formation of ATP from ADP + Pi.

Thylakoid interior
High concentration of H+
(low pH) Electron transport chain ATP synthesis

H+

H+ H+ H+
H+ H+ H+ H+ H+ H+
2 H+ H+
H2O 1 O2 H+ H+
2 NADP ATP
H+ H+ H+
H+ H+ H+ reductase H+
synthase
H+ H+ H+
e–
PC
e–
e–
PQ e–
Thylakoid
e– I Cyt
membrane

e–

Fd

Photon
Photon Photosystem II Photosystem I
H+
H+
H+
Protons are actively transported to NADP+ NADPH
interior of thylakoid compartment by +
electrons from photosystem II. H+

ATP synthase couples the formation of

ADP + Pi
ATP
Stroma
ATP to the passive diffusion of protons
Low concentration of H+
across the membrane.
(high pH)
H+
154 CHAPTER EIGHT

tons (H+) across the thylakoid membrane, which results in a
proton gradient across the membrane.
The electron carriers in the thylakoid membranes are ori-
ented so that protons move from the stroma into the interior
of the thylakoid. The interior compartment becomes acidic
with respect to the stroma. When there is sufficient light, the
ratio of H+ inside versus outside a thylakoid is usually
10,000:1, which is a difference of 4 pH units. This difference
rapidly killed the cells, extracted their carbohydrates, and
separated the different compounds from one another by pa-
per chromatography. Many compounds, including mono-
saccharides and amino acids, contained 14C (Figure 8.12).
EXPERIMENT
Question: What is the pathway of CO2 fixation in photosynthesis?

leads to the diffusion of H+ back out of the thylakoid interior

METHOD Bright light source
through specific protein channels in the thylakoid membrane. (energy for photosynthesis)
These channels are enzymes—ATP synthases—that couple
14 CO was
the diffusion of protons to the formation of ATP, just as in mi- 2
injected here.
tochondria (see Figure 7.12). In the chloroplast, the ATP is
generated in the stroma, where it will be available to provide
Thin flask of
the energy for the fixation of CO2 in the production of car- green algae
bohydrate by the Calvin-Benson cycle.

Making Carbohydrate from CO2 :

The Calvin–Benson Cycle Algae were rapidly killed and their metabolites partially
extracted by putting the cells in boiling ethanol.
At the start of this chapter we identified two distinct metabolic
pathways operating in photosynthesis. We have now discussed First
The plant extract was
run
the first pathway: the light reactions, which use light energy to spotted here and run
in two directions to
produce ATP and NADPH + H+ in the chloroplasts of green
separate compounds
plants. The second pathway, the Calvin–Benson cycle, uses this Second
from one another.
run
ATP and NADPH + H+ to incorporate CO2 into carbohydrates.
Paper
Most of the enzymes that catalyze the reactions of the chromatogram
Calvin–Benson cycle are dissolved in the chloroplast stroma
(the “soup” outside the thylakoids), and that is where those
reactions take place. However, these enzymes use the energy
in ATP and NADPH, produced in the thylakoids by the light After separation, the chromatogram was overlaid with
reactions, to reduce CO2 to carbohydrates. Because there is X-ray film that the radiation “exposed.” Each dark spot
is a compound labeled with 14C.
no stockpiling of these energy-rich coenzymes, these
Calvin–Benson cycle reactions take place only in the light, RESULTS
when these coenzymes are being generated.
GLUT
ALA
GLY SER
Isotope labeling experiments revealed the steps ASP CIT

of the Calvin–Benson cycle SUC G3P

To identify the sequence of reactions by which CO2 ends up 3PG 3PG

in carbohydrates, it was necessary to label CO2 so that it

HEXOSE-P
could be followed after being taken up by a photosynthetic
cell. In the 1950s, Melvin Calvin, Andrew Benson, and their A chromatogram made A chromatogram made
colleagues used radioactively labeled CO2 in which some of after 3 seconds of exposure after 30 seconds of
the carbon atoms were not the normal 12C, but its radioiso- to 14 CO2 shows 14 C only exposure to 14 CO2 shows
14 C in many molecules.
in 3PG.
tope 14C. Although 14C is distinguished by its emission of ra-
diation, chemically it behaves virtually identically to nonra- Conclusion: The initial product Conclusion: The carbon from CO2
dioactive 12C. In general, enzymes do not distinguish of CO2 fixation is 3PG. ends up in many molecules.
between isotopes of an element in their substrates, so 14CO2 is
treated the same way by photosynthesizing cells as 12CO2. 8.12 Tracing the Pathway of CO2 The historical photo-
graph at the top shows the apparatus Calvin and his col-
Calvin and his colleagues exposed cultures of the unicel- leagues used to follow labeled carbon dioxide molecules
lular green alga Chlorella to 14CO2 for 30 seconds. They then (14CO2) as they were transformed by photosynthesis.
 PHOTOSYNTHESIS: ENERGY FROM THE SUN 155

However, if they stopped the exposure after just 3 seconds, mediate six-carbon compound, which quickly breaks down
only one compound was labeled—a three-carbon sugar and forms two three-carbon molecules of 3PG (as Calvin and
phosphate called 3-phosphoglycerate (3PG): colleagues observed; Figure 8.14). The enzyme that catalyzes
this fixation reaction, ribulose bisphosphate carboxylase/
C OO– Carboxyl group oxygenase (rubisco), is the most abundant protein in the
world, comprising about 20 percent of all the protein in every
H C OH
plant leaf.
H C O P

H
3-Phosphoglycerate (3PG)
By tracing the steps in this manner, they soon discovered a cy-
cle that “fixes” CO2 in a larger molecule, produces a carbohy-
drate, and regenerates the initial CO2 acceptor. This cycle was
appropriately named the Calvin–Benson cycle (Figure 8.13).
The initial reaction in the Calvin–Benson cycle adds the
one-carbon CO2 to a receptor, the five-carbon compound
ribulose 1,5-bisphosphate (RuBP). The product is an inter-
ELECTRON
TRANSPORT
The Calvin–Benson cycle is made up of three processes
The Calvin–Benson cycle uses the high-energy coenzymes
made in the thylakoids during the light reactions (ATP and
NADPH) to reduce CO2 to a carbohydrate. There are three
processes that make up the cycle:

Fixation of CO2. As we saw, this reaction is catalyzed by
rubisco, and its product is 3PG.

Reduction of 3PG to form a carbohydrate, glyceralde-
hyde 3-phosphate (G3P). This series of reactions
involves a phosphorylation (using the ATP made in the
light reactions) and a reduction (using the NADPH
made in the light reactions).

ATP NADPH
CYCLE CYCLE
START 8.13 The Calvin–Benson
Cycle The Calvin–Benson
CALVIN–
BENSON 1 CO2 combines with cycle uses CO2 and the ATP
CYCLE
its acceptor, RuBP, and NADPH + H+ generat-
forming 3PG. ed in the light reactions to pro-
6 CO2
duce glucose. This diagram shows
only the key steps; the values
given are those necessary to make
5 RuMP is converted to RuBP in a reaction one molecule of glucose, which
requiring ATP. RuBP is ready to accept requires six “turns” of the cycle.
another CO2. P C C C C C P
6 RuBP
C C C P

6 ADP Carbon 12 3PG

fixation 12 ATP
6 ATP
CALVIN–BENSON CYCLE 12 ADP
6 RuMP 2 3PG is reduced to G3P
Regeneration Reduction and in a two-step reaction
of RuBP sugar production requiring ATP and
NADPH + H+.
P C C C P

12 NADPH + 12 H+

C C C P C C C P 12 NADP+
12 Pi
10 G3P 12 G3P

4 The remaining five-sixths of 2 G3P

the G3P is processed in the
complex reactions that 3 About one-sixth of the G3P is used to
produce RuMP. make sugars—the output of the cycle.
Sugars

Other carbon compounds

156 CHAPTER EIGHT

1 The fate of the carbon 2 The enzyme rubisco 3 The reaction intermediate
atom in CO2 is followed catalyzes the reaction splits into two molecules of
in red. of CO2 with RuBP. 3-phosphoglycerate (3PG).

CH2O P C

C O C C CH2O P COO –
Rubisco
C O2 + H C OH C HO C H + H C OH

Carbon H C OH C C OO– CH2O P

dioxide
CH2O P C

Ribulose 1,5- Six-carbon

bisphosphate skeleton carbohydrate generated in the cycle represent the total en-
(RuBP) of reaction
intermediate ergy yield from the harvesting of light by photosynthetic or-
ganisms. Most of this stored energy is released by glycolysis
8.14 RuBP Is the Carbon Dioxide Acceptor CO2 is added to a five- and cellular respiration and used to support plant growth,
carbon compound, RuBP. The resulting six-carbon compound imme- development, and reproduction. Much plant matter ends up
diately splits into two molecules of 3PG.
being consumed by animals, supplying them with both raw
materials and energy sources. Glycolysis and cellular respi-
ration in the animals release free energy from the plant mat-
ter for use in the animal cells.

Regeneration of the CO2 acceptor, RuBP. Most of the G3P
ends up as RuMP (ribulose monophosphate), and ATP is
used to convert this compound to RuBP. So for every Photorespiration and Its Consequences
“turn” of the cycle, with one CO2 fixed, the CO2 acceptor The enzyme rubisco, used by the Calvin-Benson cycle to fix
is regenerated. CO2 during photosynthesis, is probably the most abundant
The end product of this cycle is glyceraldehyde 3-phosphate enzyme on the planet. Its properties are remarkably identi-
(G3P), which is a three-carbon sugar phosphate, also called cal in all photosynthetic organisms, from bacteria to flower-
triose phosphate: ing plants. However, rubisco has properties that severely
limit its efficiency under certain conditions. In the discussion
H
that follows, we will identify and explore some of these lim-
C O itations and see how evolution has constructed metabolic by-
H C OH
passes around them. First we’ll look at photorespiration, a
process in which rubisco reacts with O2 instead of CO2, low-
H C O P ering the overall rate of CO2 fixation. Then we’ll examine
H some biochemical pathways and features of plant anatomy
Glyceraldehyde 3-phosphate (G3P) that compensate for the limitations of rubisco.

In a typical leaf, there are two fates for the G3P:

One-third of it ends up in the polysaccharide starch,
which is stored in the chloroplast.

Two-thirds of it is converted in the cytosol to the disac-
charide sucrose, which is transported out of the leaf to
other organs in the plant, where it is hydrolyzed to its
constituent monosaccharides: glucose and fructose.
The G3P produced in photosynthesis is subsequently used
by the plant to make other compounds. Its carbon is thus in-
corporated into amino acids, lipids, and the building blocks
of the nucleic acids.
The products of the Calvin–Benson cycle are of crucial im-
portance to the entire biosphere, for the covalent bonds of the
Rubisco catalyzes RuBP reaction with O2 as well as CO2
As its full name indicates, rubisco is a carboxylase (adding
CO2 to the acceptor molecule RuBP) as well as an oxygenase
(adding O2 to RuBP). These two reactions compete with each
other. So when RuBP reacts with O2, it cannot react with CO2.
This reaction reduces the overall CO2 that is converted to car-
bohydrates, and therefore limits plant growth.
When O2 is added to RuBP, one of the products is a two-
carbon compound, phosphoglycolate:
RuBP + O2 → phosphoglycolate + 3PG
Plants have evolved a metabolic pathway that partially re-
covers the carbon that has been channeled away from the
 PHOTOSYNTHESIS: ENERGY FROM THE SUN 157

1 In the chloroplasts,
On a hot, dry day, the stomata that allow water to evap-
RuBP reacts with O2. orate from the leaf close to prevent water loss (see Figure
Glycolate is formed. 2 Glycolate diffuses into a peroxisome, 8.1). But this also prevents gases from entering and leaving
where it is converted to glycine.
the leaf. The CO2 concentration in the leaf falls because CO2
is being used up by the light-driven photosynthetic reac-
tions, and the O2 concentration rises because of these same
reactions. As the ratio of CO2 to O2 in the leaf falls, the re-
action of rubisco with O2 is favored, and photorespiration
proceeds.

C4 plants can bypass photorespiration

In plants such as roses, wheat, and rice, the mesophyll cells,
which lie just below the surface of the leaf, are full of chloro-
plasts that contain abundant rubisco (Figure 8.16a). On a hot
day, these leaves close their stomata to conserve water. The
3 Glycine is converted to glycerate in
level of CO2 in the air spaces of the leaves falls, and that of
the mitochondria and CO2 is released.
O2 continues to rise, as photosynthesis goes on. As we have
8.15 Organelles of Photorespiration The reactions of photorespi-
ration take place in the chloroplasts, peroxisomes, and, finally, in the
mitochondria.
(a) Arrangement of cells in a C3 leaf
Upper epidermis

These cells have rubisco

Calvin–Benson cycle. The phosphoglycolate forms glycerate, and fix CO2 to RuBP to
which diffuses into membrane-enclosed organelles called form 3PG.
peroxisomes (Figure 8.15). There, a series of reactions con-
Vein
verts it to the amino acid glycine:
These cells have few
glycolate → → glycine
chloroplasts and no rubisco;
they do not fix CO2.
The glycine then diffuses into a mitochondrion, where two
glycine molecules are converted to glycerate (a three-carbon
Spongy mesophyll cell
molecule), and CO2:

2 glycine → glycerate + CO2 Stoma Lower epidermis

This pathway is called photorespiration because it consumes (b) Arrangement of cells in a C4 leaf
O2 and releases CO2. It uses ATP and NADPH produced in the
light reactions, just like the Calvin–Benson cycle. The net effect Mesophyll cells have PEP
carboxylase for the reaction of
is to take two two-carbon molecules and make one three- CO2 and PEP to form a 4-carbon
carbon molecule. So one carbon of the four is released as CO2 molecule.
and three of the carbons (75%) are recovered as fixed carbon.
In other words, photorespiration reduces net carbon fixation
Bundle sheath cells have rubisco
by 25 percent compared with the Calvin–Benson cycle.
for the reaction of RuBP with CO2
How does rubisco “decide” whether to act as an oxyge- released from the 4-carbon
nase or a carboxylase? First, rubisco has 10 times more affin- compound.
ity for CO2 than O2, and so favors CO2 fixation. Another con-
sideration is the relative concentrations of CO2 and O2 in the Close association permits CO2 pumping
leaf. If O2 is relatively abundant, rubisco acts as an oxyge- from mesophyll cells to bundle sheath
nase, and photorespiration ensues. If CO2 predominates, ru- cells for the Calvin–Benson cycle.

bisco fixes it, and the Calvin–Benson cycle occurs. Tempera-

8.16 Leaf Anatomy of C3 and C4 Plants Carbon dioxide
ture is also a factor: photorespiration is more likely at high fixation occurs in different organelles and cells of the leaves
temperatures. in (a) C3 and (b) C4 plants.
158 CHAPTER EIGHT

(a)
1 PEP carboxylase in C4 mesophyll cells 2 Oxaloacetate diffuses through plasmo-
catalyzes the formation of the 4-carbon desmata to a bundle sheath cell, where
compound oxaloacetate. it is decarboxylated, releasing CO2.

Bundle
sheath cell
Mesophyll
cell
3 Starch grains in the
bundle sheath cell
indicate that the
Calvin–Benson cycle
is active and that
glucose (and then
starch) is being
produced.

8.17 The Anatomy and Biochemistry of C4 Carbon Fixation

Carbon dioxide is fixed initially in the mesophyll cells, but enters the
Calvin–Benson cycle in the bundle sheath cells.
Mesophyll (b) There is an interconnected biochemical pathway for CO2 assimila-
cell tion between the two cell types.

(b) Plasma
membrane
seen, rubisco acts as an oxygenase, and photorespiration oc-
Mesophyll curs, under these conditions. Because the first product of CO2
Cell wall
cell CO2
fixation in these plants is the three-carbon molecule 3PG, they
PEP
are called C3 plants.
Corn, sugarcane, and other tropical grasses (Figure 8.16b)
Carboxylation Regeneration also close their stomata on a hot day, but their rate of pho-
tosynthesis does not fall, nor does photorespiration occur.
C4 cycle
They keep the ratio of CO2 to O2 around rubisco high so that
4C compound 3C compound rubisco continues to act as a carboxylase. They do this in
part by making a four-carbon compound, oxaloacetate, as the
first product of CO2 fixation, and so are called C4 plants.
Bundle C4 plants perform the normal Calvin–Benson cycle, but
sheath Decarboxylation 3C compound they have an additional early reaction that fixes CO2 without
cell
losing carbon to photorespiration, greatly increasing the over-
all photosynthetic yield. Because this initial CO2 fixation step
CO2
can function even at low levels of CO2 and high tempera-
5C sugar tures, C4 plants very effectively optimize photosynthesis un-
Carboxylation der conditions that inhibit it in C3 plants.
Regeneration
Calvin–Benson C4 plants have two separate enzymes for CO2 fixation,
cycle 3C sugar located in two different parts of the leaf (Figure 8.17). One
Triose-P
enzyme, present in the cytosol of mesophyll cells near the
Reduction
surface of the leaf, fixes CO2 to a three-carbon acceptor com-
pound, phosphoenolpyruvate (PEP), to produce the four-
carbon fixation product, oxaloacetate. This enzyme, PEP car-
boxylase, has two advantages over rubisco:

It does not have oxygenase activity.

It fixes CO2 even at very low CO2 levels.

8.1 Comparison of Photosynthesis in C3 and C4 Plants
VARIABLE C3 PLANTS
Photorespiration Extensive
Perform Calvin–Benson cycle? Yes
Primary CO2 acceptor RuBP
CO2-fixing enzyme Rubisco (RuBP carboxylase/oxygenase)
First product of CO2 fixation 3PG (3-carbon compound)
Affinity of carboxylase for CO2 Moderate
Photosynthetic cells of leaf Mesophyll
Classes of chloroplasts One
So even on a hot day when the stomata are closed, the CO2
concentration in the leaf is low, and the O2 concentration is
high, PEP carboxylase just keeps on fixing CO2.
Oxaloacetate diffuses out of the mesophyll cells and
through plasmodesmata into the bundle sheath cells, located in
the interior of the leaf. The chloroplasts in bundle sheath cells
contain abundant rubisco. There, the four-carbon oxaloacetate
loses one carbon, forming CO2 and regenerating the three-
carbon acceptor compound, PEP, in the mesophyll cells. Thus,
the role of PEP is to bind CO2 from the air in the leaf and carry
it to the bundle sheath cells, where it is “dropped off” at ru-
bisco. This process essentially pumps up the CO2 concentra-
tion around rubisco, so that it acts as a carboxylase and begins
the Calvin–Benson cycle.
Kentucky bluegrass, a C3 plant, thrives on lawns in April
and May. But in the heat of summer, it does not do as well,
and crabgrass, a C4 plant, takes over the lawn. The same is
true on a global scale for crops: C3 plants, such as soybeans,
rice, wheat, and barley, have been adapted for human food
production in temperate climates, while C4 plants, such as
corn and sugarcane, originated and are grown in the tropics.
Table 8.1 compares C3 and C4 photosynthesis.
C3 plants are certainly more ancient than C4 plants. While
C3 photosynthesis appears to have begun about 3.5 billion
years ago, C4 plants appeared about 12 million years ago. A
possible factor in the emergence of the C4 pathway is the de-
cline in atmospheric CO2. When dinosaurs ruled Earth 100
million years ago, the concentration of CO2 in the atmosphere
was four times what it is now. As CO2 levels declined there-
after, the more efficient C4 plants would have had an advan-
tage over their C3 counterparts.
CAM plants also use PEP carboxylase
Other plants besides the C4 species use PEP carboxylase to fix
and accumulate CO2. Such plants include some water-storing
PHOTOSYNTHESIS: ENERGY FROM THE SUN 159
C4 PLANTS
Minimal
Yes
PEP
PEP carboxylase and rubisco
Oxaloacetate (4-carbon compound)
High
Mesophyll + bundle sheath
Two
plants (called succulents) of the family Crassulaceae, many
cacti, pineapples, and several other kinds of flowering plants.
The CO2 metabolism of these plants is called crassulacean
acid metabolism, or CAM, after the family of succulents in
which it was discovered. CAM is much like the metabolism
of C4 plants in that CO2 is initially fixed into a four-carbon
compound. In CAM plants, however, the processes of initial
CO2 fixation and the Calvin–Benson cycle are separated in
time, rather than in space.

At night, when it is cooler and water loss in minimized,
the stomata open. CO2 is fixed in mesophyll cells to form
the four-carbon compound oxaloacetate, which is con-
verted to malic acid.

During the day, the accumulated malic acid is shipped to
the chloroplasts, where decarboxylation supplies the CO2
for operation of the Calvin–Benson cycle, and the light
reactions supply the necessary ATP and NADPH + H+.
Metabolic Pathways in Plants
Green plants are autotrophs and can synthesize all the mol-
ecules they need from simple starting materials: CO2, H2O,
phosphate, sulfate, and ammonium ions (NH4+). NH4+ is
needed for amino acids and comes either from the conver-
sion of nitrogen-containing molecules in soil water taken up
by the plant’s roots, or from the bacterial conversion of N2
gas from the atmosphere.
The light reactions of photosynthesis generate ATP and
NADPH, which are used to synthesize carbohydrates. These
compounds can then be used in cellular respiration to pro-
vide energy for processes such as active transport and an-
abolism. Both cellular respiration and fermentation can oc-
cur in plants, although the former is far more common. Plant
cellular respiration, unlike photosynthesis, takes place both
in the light and in the dark. Because glycolysis occurs in the
160 CHAPTER EIGHT
(triglycerides)
CO2
Glycerol
3PG
RuBP
CALVIN–BENSON
CYCLE Pyruvate
RuMP G3P
GLYCOLYSIS
Nucleotides
Polysaccharides
Hexose-P
Nucleic acids
8.18 Metabolic Interactions in a Plant Cell The products of the
Calvin–Benson cycle are used in the reactions of cellular respiration
(glycolysis and the citric acid cycle).
cytosol, respiration in the mitochondria, and photosynthe-
sis in the chloroplasts, all these processes can proceed si-
multaneously.
Photosynthesis and respiration are closely linked through
the Calvin–Benson cycle (Figure 8.18). The partitioning of
G3P is particularly important:

Some G3P from the Calvin–Benson cycle can be convert-
ed to pyruvate, the end product of glycolysis. This pyru-
vate can be used in cellular respiration for energy, or its
carbon skeletons can be used anabolically to make lipids,
proteins, and other carbohydrates (see Figure 7.17).

Some G3P can enter a pathway that is the reverse of gly-
colysis (the gluconeogenic pathway). In this case,
sucrose is formed and transported to the nonphotosyn-
thetic tissues of the plant, such as the root.
Energy flows from sunlight to reduced carbon in photo-
synthesis to ATP in respiration. Energy can also be stored in
the bonds of macromolecules such as polysaccharides, lipids,
and proteins. For a plant to grow, energy storage (as body
structures) must exceed energy release; that is, overall carbon
fixation by photosynthesis must exceed respiration. This
principle is the basis of the ecological food chain, as we will
see in later chapters.
Nitrogen
from soil
Lipids
NH+4
Fatty acids α-Ketoglutarate Amino acids
CITRIC ACID
Acetyl CoA CYCLE Proteins
(starch)
Chapter Summary
Identifying Photosynthetic Reactants and Products

Photosynthesizing plants take in CO2, water, and light
energy, producing O2 and carbohydrates. The overall reaction is
6 CO2 + 12 H2O + light → C6H12O6 + 6 O2 + 6 H2O

The oxygen atoms in the O2 produced by photosynthesis
come from water, not from CO2. Review Figures 8.1, 8.2.
See Web/CD Tutorial 8.1
The Two Pathways of Photosynthesis: An Overview

In plants, photosynthesis takes place in chloroplasts.

In the light reactions of photosynthesis, electron transport
and photophosphorylation produce ATP and reduce NADP+ to
NADPH + H+. Review Figure 8.3
+

ATP and NADPH + H are needed for the reactions that fix
and reduce CO2 in the Calvin–Benson cycle, forming carbohy-
drates. Review Figure 8.3
The Interactions of Light and Pigments

Light energy comes in packets called photons, but it also has
wavelike properties.

Absorption of a photon puts a pigment molecule in an excit-
ed state that has more energy than its ground state. Review
Figure 8.4

Pigments absorb light in the visible spectrum. Review
Figure 8.5

Each compound has a characteristic absorption spectrum. An
action spectrum reveals the biological effectiveness of different
wavelengths of light. The absorption spectrum of the plant pig-
ment chlorophyll a correlates well with the action spectrum for
photosynthesis. Review Figures 8.6
 PHOTOSYNTHESIS: ENERGY FROM THE SUN 161

Chlorophylls and accessory pigments form antenna systems

for absorption of light energy. Review Figure 8.7

An excited pigment molecule may lose its energy by fluores-
cence or by transferring it to another pigment molecule. Review
Figure 8.8
Electron Transport, Reductions, and
Photophosphorylation

Noncyclic electron transport uses two photosystems (I and II)
and produces ATP, NADPH + H+, and O2. Photosystem II uses
P680 chlorophyll, from which light-excited electrons are passed
to a redox chain that drives chemiosmotic ATP production.
Light-driven oxidation of water releases O2 and passes electrons
from water to the P680 chlorophyll. Photosystem I passes elec-
trons from P700 chlorophyll to another redox chain and then to
NADP+, forming NADPH + H+. Review Figure 8.9

Cyclic electron transport uses P700 chlorophyll and produces
only ATP. Its operation maintains the proper balance of ATP and
NADPH + H+ in the chloroplast. Review Figure 8.10

Chemiosmosis is the mechanism of ATP production in pho-
tophosphorylation. Electron transport pumps protons from the
stroma into the thylakoids. Diffusion of the protons back to the
stroma via ATP synthase channels drives ATP formation.
Review Figure 8.11. See Web/CD Tutorial 8.2
Making Carbohydrate from CO2: The Calvin–Benson Cycle

The Calvin–Benson cycle makes sugar from CO2. This path-
way was elucidated through the use of radioactive tracers.
Review Figure 8.12. See Web/CD Tutorial 8.3

The Calvin–Benson cycle consists of three phases: fixation of
CO2, reduction and carbohydrate production, and regeneration
of RuBP. RuBP is the initial CO2 acceptor, and 3PG is the first
stable product of CO2 fixation. The enzyme rubisco catalyzes
the reaction of CO2 and RuBP to form 3PG. Review Figures
8.13, 8.14. See Web/CD Activity 8.1
Photorespiration and Its Consequences

The enzyme rubisco can catalyze a reaction between O2 and
RuBP in addition to the reaction between CO2 and RuBP. This
reaction with O2 is called photorespiration and significantly re-
duces the efficiency of photosynthesis. The reactions that consti-
tute photorespiration are distributed over three organelles:
chloroplasts, peroxisomes, and mitochondria. Review Figure
8.15

At high temperatures and low CO2 concentrations, the oxyge-
nase function of rubisco is favored.

C4 plants bypass photorespiration with special chemical reac-
tions and specialized leaf anatomy. In C4 plants, PEP carboxy-
lase in chloroplasts of the mesophyll cells initially fixes CO2 in a
four-carbon compound, which then diffuses into bundle sheath
cells, where its decarboxylation produces locally high concentra-
tions of CO2. Review Figure 8.16, 8.17. See Web/CD Activity 8.2

CAM plants operate much like C4 plants, but their initial CO2
fixation by PEP carboxylase is temporally separated from the
Calvin–Benson cycle, rather than spatially separated as in C4
plants.
Metabolic Pathways in Plants

Photosynthesis and respiration are linked through the
Calvin–Benson cycle, the citric acid cycle, and glycolysis.
Review Figure 8.18

To survive, a plant must photosynthesize more than it respires.
Self-Quiz
1. In noncyclic photosynthetic electron transport, water is
used to
a. excite chlorophyll.
b. hydrolyze ATP.
c. reduce chlorophyll.
d. oxidize NADPH.
e. synthesize chlorophyll.
2. Which statement about light is true?
a. An absorption spectrum is a plot of biological effective-
ness versus wavelength.
b. An absorption spectrum may be a good means of identi-
fying a pigment.
c. Light need not be absorbed to produce a biological effect.
d. A given kind of molecule can occupy any energy level.
e. A pigment loses energy as it absorbs a photon.
3. Which statement about chlorophylls is not true?
a. They absorb light near both ends of the visible spectrum.
b. They can accept energy from other pigments, such as
carotenoids.
c. Excited chlorophyll can either reduce another substance
or fluoresce.
d. Excited chlorophyll may be an oxidizing agent.
e. They contain magnesium.
4. In cyclic electron transport,
a. oxygen gas is released.
b. ATP is formed.
c. water donates electrons and protons.
d. NADPH + H+ forms.
e. CO2 reacts with RuBP.
5. Which of the following does not happen in noncyclic elec-
tron transport?
a. Oxygen gas is released.
b. ATP forms.
c. Water donates electrons and protons.
d. NADPH + H+ forms.
e. CO2 reacts with RuBP.
6. In the chloroplasts,
a. light leads to the pumping of protons out of the thylakoids.
b. ATP forms when protons are pumped into the thylakoids.
c. light causes the stroma to become more basic than the
thylakoids.
d. protons return passively to the stroma through protein
channels.
e. proton pumping requires ATP.
7. Which statement about the Calvin–Benson cycle is not true?
a. CO2 reacts with RuBP to form 3PG.
b. RuBP forms by the metabolism of 3PG.
c. ATP and NADPH + H+ form when 3PG is reduced.
d. The concentration of 3PG rises if the light is switched off.
e. Rubisco catalyzes the reaction of CO2 and RuBP.
8. In C4 photosynthesis,
a. 3PG is the first product of CO2 fixation.
b. rubisco catalyzes the first step in the pathway.
c. four-carbon acids are formed by PEP carboxylase in
bundle sheath cells.
d. photosynthesis continues at lower CO2 levels than in
C3 plants.
e. CO2 released from RuBP is transferred to PEP.
162 CHAPTER EIGHT
9. Photosynthesis in green plants occurs only during the day.
Respiration in plants occurs
a. only at night.
b. only when there is enough ATP.
c. only during the day.
d. all the time.
e. in the chloroplast after photosynthesis.
10. Photorespiration
a. takes place only in C4 plants.
b. includes reactions carried out in peroxisomes.
c. increases the yield of photosynthesis.
d. is catalyzed by PEP carboxylase.
e. is independent of light intensity.
For Discussion
1. Both photosynthetic electron transport and the Calvin–
Benson cycle stop in the dark. Which specific reaction stops
first? Which stops next? Continue answering the question
“Which stops next?” until you have explained why both path-
ways have stopped.
2. In what principal ways are the reactions of electron transport
in photosynthesis similar to the respiratory chain and oxida-
tive phosphorylation discussed in Chapter 7? Differentiate
between cyclic and noncyclic electron transport in terms of (1)
the products and (2) the source of electrons for the reduction
of oxidized chlorophyll.
3. The development of what two experimental techniques made
it possible to elucidate the Calvin–Benson cycle? How were
these techniques used in the investigation?
4. If water labeled with 18O is added to a suspension of photo-
synthesizing chloroplasts, which of the following compounds
will first become labeled with 18O: ATP, NADPH, O2, or 3PG?
If water labeled with 3H is added to a suspension of photo-
synthesizing chloroplasts, which of the same compounds will
first become radioactive? If CO2 labeled with 14C is added to a
suspension of photosynthesizing chloroplasts, which of those
compounds will first become radioactive?
5. The Viking lander was sent to Mars in 1976 to detect signs of
life. Explain the rationale behind the following experiments
this unmanned probe performed:
a. A scoop of dirt was inserted into a container and 14CO2 was
added. After a while during the Martian day, the 14CO2 was
removed and the dirt was heated to high temperatures.
Scientists monitoring the experiment back on Earth looked
for the release of 14CO2 as a sign of life.
b. The experiment in (a) was performed, except that the dirt
was heated to high temperature for 30 minutes and then
allowed to cool to Martian temperature right after scooping
and before the 14CO2 was added. If experiment (a) released
14
CO2, then this experiment should not release it, if living
things were present.